A New Genus and Species of Nine-Primaried Oscine of Uncertain Affinities from Peru

THE AUK A QUARTERLY JOURNAL OF ORNITHOLOGY

VoL. 93 J•JL¾1976 No. 3

A NEW GENUS AND SPECIES OF NINE-PRIMARIED OSCINE OF UNCERTAIN AFFINITIES FROM PERU

G•ORG• H. LoweRY, JR., ^•r• D^• A. T^LLM^•

T• frequencywith which hitherto unknown genera and speciesof birds have continued to turn up in Peru in recent years is indeed remarkable. Among the noveltiesdiscovered and duly describedsince 1964 are 2 new generaand 10 new species•, someof which have been spectacular(for example, the tanagers Buthraupis aureodorsalisand Wetmorethraupissterrhopteron). Also now in preparationby personnel here at this .museumand by avian systematistsat other ornithological centers are descriptionsof at least 6 additional new species(an owl, 2 hummingbirds,2 flycatchers,and a wren). This paper describesstill another recent Peruvian discovery, a bird that we call the "Pardusco," becausethat is the name applied to it by our Peruvian field assistantswho live near the region where it is now known to occur. An adult male and an immature male were obtained in June 1973 and 8 additional specimens(5 males and 3 females) were taken in June 1974, by our resident assistant Manuel Villar, while a memberof our field parties of those years. And, finally, in January, June, and July 1975, the remainingspecimens, 22 males and 15 females, were procuredby another one of our field parties, composedof Villar, Robert S. Kennedy, Carol S. O'Neill, Theodore Parker, III, and Reyes Rivera A. All encounters with the bird have been in the isolated and semi-isolatedwooded tracts of low trees and shrubs (elfin forest) near the crest of the Carpish Ridge of the eastern cordillera of the Andes in the Departamento de Hu&nuco, above Acomayo. It is the region in

•Synallaxis courseni Blake, Percnostola macrolopha Berlioz, Grailaria eludens Lowery and O'Neill, Conioptilon mcilhennyi Lowery and O'Neill, tlemispingusparodi• Weske and Terborgh, Hemispingusrufosuperciliarls Blake and Hocking, Buthraupls aureodorsalisBlake and Hocking, Wetmorethraupis sterrhopteron Lowery and O•Neill, Cacicus koepckeae Lowery and O'Neill, and Agelaius xanthophthalmus Short.

415 The Auk 93: 415-428. July 1976 P•a•DreSCO,Nephelornis oneilli A NEW OENI, J$ AND $I•ECI1•$ rRO• I•]•RU From a watercolor painting by John P. O'Neill 416 Low•a¾ ^up T^LLM^N [Auk, Vol. 93 which Buthraupis aureodorsalisand Hemispingusrufosuperciliaris were discoveredand is still the only location from which they are known. These same woodlandsare inhabited by Ampelion [Doliornis] sclateri, a speciesof cotingaknown until 3 yearsago only from 2 old spedmens from the Departamento de Junin but now found to be fairly common above Acomayo in suitable habitats. That this regionhas yieldedsuch unexpected discoveries is surprising, for it lies in a part of Peru that must be consideredone of the better known sectionsof the republic. In 1922 and 1923 John T. Zimmer passedthrough or within easy striking distanceof the forest that these speciesfrequent (see Zimmer 1930), and our own field parties in the 1960'sand early 1970'slikewise often operatedwithin sight of the same forest. One can only speculatehow many more such ornithological treasure-trovesawait discoveryin the ruggedmassifs of the Peruvian Andes. From the outset,we have consideredthe Parduscoto be a bird of uncertain familial relationships.For reasonsstated beyond, we cannot affiliate it with any known genusof tanager, honeycreeper,or finch. Becauseof certain morphologicalpeculiarities, mainly ones pointed out to us by Walter J. Bock followinghis study of a skeletonand a wet- preservedspedmen we were able to supplyhim, we feel compelledto erect for it a new genusand to leave its familial placementto the outcome of further studies.

Nephelornis gen. nov. TYPz-sPzcms: Nephelornis oneJill Lowery and Tallman. DIAGNOSIS:A small, nine-primaried oscinewith the wing formula 6 > 7 > 5 > 4 > 8 > 3 > 2 > 1 > 9; wing longer than tail, with the ratio approximately 1.16:1.00; tail slightly graduated,with the central rectricesthe longestby approximately 6 to 8 ram; shapeof rectricesmoderately broad and terminallyrounded; culmen distinctly arched; bill small in overall size and approximately half as deep and as wide at the base as it is long; tomium decurved and with a barely perceptiblerhamphothecal subterminalindentation that would hardly qualify as a notch; nostrils fully exposed, roughly oval in shape,and with the superiorhemisphere of each overhungby a membrane; rictal bristlesrather long but weak; a few bristleson the chin and face; toes and the laminiplantar tarsometatarsusconspicuously strong; middle toe (no. III) longest and length without claw more than half that of the tarsometatarsus; toe no. IV without claw slightly longer than toe no. II without claw and both shorter than hind toe (no. I); hind claw fully twice the size and length of the other claws; skull fairly typical of nine-primaried oscines(Fig. 1A and B) with no diagnostic palatal or mandibular features but in general shape and proportions readily separable from the skull of any genus with which it has been compared, includingHemispingus, Chlorospingus, Xenodacnis, Diglossa, Conirostrum, Iridophanes, and a wide array of emberizine finches; the tongue slightly frilled with the lateral edges raised to form a shallow groove (Fig. 2A); the basihyale flattened, as is July 1976] The Pardusco 417

Fig. 1. Skull of Nephelornis oneilli LSUMZ 80122). (A) Lateral view with mandible in position. (B) Ventral view without mandible. (C) Lateral view of basihyale and proximal end of ceratobranchiale.(D) Schematiccross section of basihyaleat its approximatemidpoint. Drawings by Dorothea Goldys. typical for nine-primariedoscines (Figs. 1C and D); a small slip of M. tracheo- lateralisinserting onto the proximal end of the ceratobranchiale,a conditionseen in many passerinebirds but lost in most nine-primaried oscines(Figs. 2B and 3A and B); M. hypoglossusanterior present,a conditionfound in many passerinesbut lost in most, if not all, nine-primariedoscines (Figs. 3A and C); the M. cerato- hyoideustaking origin partly from the medial surface of the ceratobranchialeas is 418 LOWERY' AND TALLMAN' [Auk, Vol. 93

tr h B hg o

Msth bm 'Msh Fig. 2. Tongue apparatusof NephelornisoneJill (LSUMZ 77650). (A) Dorsal view of the corneoustongue to the frilled tip and slightly upturned lateral edges. (B) Ventral view of the entire tonguemusculature in placein the head; the iV[ s h and M m h have been removed on the right side. Abbreviations: branchiomandibularis;M c g : M. ceratoglossus;M c h : iV[. ceratohyoideus; M g g •-• iV[. genioglcssus;M hg a = iV[. hypoglossusanterior; iV[ hg o hypoglossusobliquus; M m h = M. mylohyoideus;M s h : iV[. serihyoideus; iV[ st h : iV[. stylohyoideus;M th h = iV[. thyreohyoideus;iV[ tr h: M. tracheo- hyoideus;M tr 1 = iV[. tracheolateralis.Drawings by Dorothea Goldys.

usual for most passerinesand for nine-primariedcscines in particular, but also taking its origin partly from the lateral surface of the ceratobranchiale,a feature that may be unique among nine-primariedoscines (Figs. 2B and 3A and B). COLO•.ATtoZV:The only spedes presently known is rather nondescript, mostly plain brown without streaks, spcts, distinct wing bars, a superciliary line, or any facial marks. July 1976] The Pardusco 419

st h

M trh hg o M tr Ix hg a

gg M thh • cg

tr h

c gg

'M hg a M hgo

Fig. 3. Tongue apparatus of Nephelornis oneilli (LSUMZ 77650). (A) Ventral view of the entire tongue musculature removed from the head and spread out to show the musclesclearly. (B) Dorsal view of the right hyoid horn showing the musclesattached to it. (C) Ventral view of the paired paraglossaliashowing the musclesattached to it. SeeFigure 2 for abbreviations.Drawings by DorotheaGoldys. 420 LOWERY AND TALLMAN [Auk, Vol. 93

Fig. 4. A Pardusco[oHowing its capturein a mist net at BosqueUnchog. Photo by R. S. Kennedy.

Sr.zc•s: No obvioussexual dimorphism but femalesaverage slightly smaller and weigh less than do males.

Nephelornis oneJill sp. nov.

P•u•usco

TYPE: Adult male (skull fully ossified);Louisiana State UniversityMuseum of Zoologyno. 81114; BosqueUrichog, on passbetween Churubamba and Hacienda Paty aboveAcomayo, 09o41 ' S, 75ø07' W, elevationapproximately $$92 meters• July 1976] The Pardusco 421

kms

Fig. 5. Map of the Acomayoarea of the Departamentode Hu•nuco•Peril, showing the locationof place-namesand physicalfeatures mentioned in the text.

Depto. Hu•nuco, Peril; 16 July 1975; collectedby TheodoreParker, III; original number 922. DmG•os•s: Same as for the genus,of which it is the only known member. DEscmPT•o• oF HOEOTYP•: General color wholly brownish; entire dorsum Mummy Brown (capitalizedcolor namesare from Ridgway 1912); primariesand secondaries Blackish Brown (3), faintly edged anteriorly with Tawny-Olive; greater secondary covertsdull brown, with Ochraceous-Tawnyterminal edgingsthat form an indistinct wing bar; upper surfaceof rectricesBlackish Brown, with outer web narrowly edged 422 Low• AND TALLMAN [Auk, Vol. 93

Fig. 6. A panoramicview of BosqueUnchog, the forestedarea on the ridge in center and left side of photograph. The woodlandsbelow the crest of the ridge is one of the habitats where Nephelornisoneilli is presentlyknown to occur. The low area at the base of the woodlands is the pass between Churubamba and Hacienda Patjr. White lines are the trails followed almost daily by LSUMZ personnelduring their visits; X, a few of the placeswhere specimensof the specieswere taken; O, some additional sites where the specieswas observed; star, the site where the holotype was obtained. Photo by R. S. Kennedy.

with Light BrownishOlive; underpartsbetween Tawny Olive and Ochraceous-Tawny, shadinginto Light Buff on the throat; flanks and undertail covertsslightly darker than the abdomen; eye-ring almost imperceptiblypaler than the face; no superciliary line or differentiation in color between the lores and the remainder of the face. (See Frontispiece and Fig. 4.) PAe•ATX•PXCVAa•A•XO•r: The type series is remarkably uniform in coloration. Only one specimen,LSUMZ 74730, an immature male in postjuvenal (first prebasic) molt that was obtained on 15 June, shows significant differences from the remainder of the series,and then only to the extent that the juvenal feathers are an appreciably darker brown throughout. Only the pectoral portion of the ventral tract appears to have been largely replaced. The remiges, rectrices,and greater secondary coverts are almost devoid of the pale edgings evident in older birds. We detect no difference in the shape or width of the rectricesin the juvenal specimenother than the possibility that these feathers may be slightly more pointed. July 1976] The Pardusco 423

Fig. 7. A close-up view of one of the wooded trac• at Bosque Unchog where Nephdornis o•illi • •mmon. Photo by R. S. K•dy.

The labels of 11 male specimensthat show the extent of ossificationof the skull indicate that in 10 the skull was fully ossified and that in one it was C•partially ossified." But the plumage of the last differs in no. way from the remainder. In 10 female specimenswhere the same information is given, all 10 carry the notation that the skull was fully ossified.Two male specimens,taken on 14 and 16 July respectively, show the size of the testesto have been "enlarged." The measurementsof the testes in both specimensare given as 8 X 5 mm. In 10 additional males where the size of the testes is stated the measurements are 3.5 X 2.5 mm or smaller. In 10 females the ovary measured 5 X 4 mm or smaller and in all cases was said to be 'Cnot enlarged? Some of the foregoing specimensare almost certainly birds of the year, but they provide no clues whereby they can be recognizedas such. Apparently, once the postjuvenal (first prebasic) molt is complete, individuals of different age groups are indistinguishableby plumage criteria. No sexual dimorphismis evident in the seriesexcept that, as already noted, males average slightly larger and weigh more than females. COLORO1• SOFT PARTS: Irises brown; maxilla dark brown or horn colored;mandible flesh-colored; tarsi light brown; toe pads yellowish. R•NOE: So far as known, near the summit of the easterncordillera of the Andes, in the Departamento de Hu•nuco, above Acomayo. SPECimEnsEXAminED: Forty-seven, including 29 males(24 skins,some with partial skeletons;3 completeskeletons; and 2 alcohol-preservedspecimens) and 17 skins and 1 alcohol-preservedspecimen of females,all from the type locality or within a few kilometersthereof at BosqueCahuincho and BosqueQuiullacocha (see Fig. 424 LOWERy ^Nn T^•M^N [Auk, Vol. 93

MEASUREMENTSIN MILLIMETERS: Males (N ---- number, next the holotype, then the mean followed by its Standard Deviation, and finally the mensural range in parentheses):wing unflattened,N26, 69.7, 67.6 ñ 2.92 (62.0-71.6); tail, N22, 59.3, 58.8 •__ 2.28 (54.8-62.5); tarsus, N26, 23.4, 23.4 ñ 0.68 (22.2-24.7); middle toe without claw, N26, 14.1, 13.6 ñ 0.61 (12.5-14.8); exposedculmen, N25, 11.0, 10.4 ___0.52 (10.0-11.2); depth of bill at base of exposed culmen, N26, 5.6, 5.2 ___0.19 (4.9-g.6); width of bill at base of culmen,N25, 5.4, 5.3 ñ 0.28 (5.0-5.8). Females (N ---- 17; first figure is the mean, then its Standard Deviation followed by the range in parentheses):wing unflattened,64.2 -+ 2.05 (61.2-69.2); tail, 56.6 ñ 2.03 (53.5-59.5); tarsus,21.9 q- 0.66 (20.0-22.8); middle toe without claw, 13.1 q- 0.47 (12.3-14.0); exposedculmen, 10.6 -+ 0.36 (10.1-11.4); depth of bill at base of culmen, 5.0 q- 0.19 (4.8-5.3); width of bill at base of culmen, 5.1 q- 0.25 (4.6-5.5). WEXCXXTS:Fifteen males averaged 17.5 ___0.94 (16.0-19.0) grams; 13 females averaged 14.9 q- 0.68 (13.5-15.5). An additional male that was prepared as a skeletonis labeledas having weighed 15.0 grams,but missexingor some other error m•y have occurred. ET¾•XOLOC¾:The name Nephelornis, "bird of clouds or mist," comes from the Greek words nephel•, cloud, and ornis, bird, and alludes to the gloomy, fogbound habitat near the crest of the Carpish Mountains where the speciesoccurs. The name is masculine in gender. We take pleasure in applying the specific epithet oneJillin honor of our colleagueJohn P. O'Neill, in recognitionof the great con- tributions he has made to Peruvian ornithologyin the past decadeand a half. In that period he has led a field party to Peru every year except one. RE•X^RnS:We are greatly indebtedto Walter J. Bock for providing us with the results of his dissectionsof the jaw and tongue musculatureof one of our wet- nreservedspecimens. He finds two charactersin particular that are consistentwith the premise that Nephelornis is a rather primitive offshoot in the radiation of the nine-primaried assemblage: (1) the presenceof M. hypoglossusanterior, which, so far as is known, is lost in other nine-prlmaried oscinesbut retained in vireos and the Olive Warbler (Peucedramustaeniatus); and (2) the fact that M. ceratohyoideus takes orlein both from the lateral surfaceof the ceratobrachiale,as well as from its medial surface. But where in this nine-primariedassemblage its closestgeneric or even familial or subfamilial relationslie remainsenigmatic.

HABITS AND BEHAVIOR Thanksto the carefulobservations of TheodoreParker, III, a member of our Peruvian field partiesin both 1974 and 1975, we have considerable information concerningthe behavior and habits of the Pardusco..In the summary that follows we have drawn freely on Parker's superb field notes. Within its limited elevationaland known geographicrange this species is surprisingly common. It occurs in the isolated and semi-isolated forestedtracts of low trees and bushesthat extend irregularly upward from continuouscloud forest, from 9800 to 11,500 feet (ca. 3000 to 3500 m). The speciesfrequents mainly the edges of these woodlands and the shrubsthat dot sphagnumbogs. See Figures 5, 6, and 7. Tree heights in the wooded areas range from 12 to 30 feet (ca. 4-10 m), July 1976] The Pardusco 425 while bushesalong the edgesand in the bogs vary from 3 to 5 feet (ca. 1.0-1.5 m) in height and 2 to 3 feet (0.6-0.9 m) in lateral spread. Both treesand busheshost a wide variety of epiphyticgrowth, especially mossesand lichens. Grazed grasslandsprinkled with ferns separatesthe isolated patches of elfin forest. The habitat of the Parduscois usually fogboundfor a greatpart of eachday. Rainsfall almostdaily, especially duringthe rainy season(November to April). Parduscohabitat, though sparselyinhabited by Quechuasin places,is for the mostpart too remote and wet for human use. Consequently,though the known geographic range of the speciesis extremelysmall, habitat destructionor other pressuresappear not to threaten its population. Parker and his companionsfound that the Parduscomoved either in conspecificflocks of 5 to 15 individualsor in mixedflocks of tanagers, conebills(Conirostrum), and flower-piercers(Diglossa). The species could be located from a distanceby the "seep" notes uttered constantly by membersof a group moving from bush to bush. To Parker they were in this regard reminiscentof the CommonBushtit (Psaltriparus minimus)of North America.The contactcall ("seep") and a soft "chip" were the only vocalizationsheard, but apparentlynone of the visits of our personnelto the regionhas yet coincidedwith the Pardusco'sbreeding season.The label of a female taken on 14 July carriesthe notation that an old broodpatch was evident, but the specimenshows no signs of postnuptial (prebasic) molt. Conspecificflocks were usually compact and flock memberswere constantlyon the move, rarely lingeringat one foragingsite. Though they werenot shy and couldbe easilyapproached, they did not respond to squeakingsounds that North American ornithologistscommonly employin callingup smallbirds. Individualswould fly into the mid- portion of a bush and then work upward and out onto the limbs. They seemedto prefer shrubsor trees with densecrowns of clustered,small leaves. They fed deliberately,gleaning lower and (less frequently) upper leaf surfacesand stems. Insects unquestionablymake up the greater part of the bird's diet, though plant or insect secretionsare perhapstaken from the undersideof leaves.The stomachsof all specimens cursorilyinspected contained numerous insects. Four preserved stomachscritically examinedcontained the remains of a spider and numerousidentifiable parts of Lepidoptera,Coleoptera, Homoptera, and Diptera, as well as a small amount of fibrous plant material. Foragingposture was usually upright though Parduscossometimes hung head downward to reach previously uninspectedsurfaces or straightenedtheir legs to reach leaves situated above them. On working up through a densely foliated bush to the top, the birds sometimes 426 Low•u¾ AND TALL3•AN [Auk, Vol. 93 perchedbriefly in an upright position, peering from side to side, and then flew to anotherbush where other individualswere already foraging. At times, while passing short distancesbetween bushes, individuals employeda slow, fluttering flight, less than a meter above the ground, reminiscent to Parker of some wrens (Cistothorus) and furnariids (Asthenes),but usually the flight was rapid and direct. In one in- stancetwo individualsof a flock descendedto the groundin a sphagnum bog,where they hoppedabout on the moss,probing and gleaningblades ef protruding grasses. Occasionallygroups of Parduscoswould join flocks of forest tanagers and honeycreepers,and at suchtimes they ascendedto the forestcanopy, but they seemeddefinitely to prefer low, edge situations.Species seen in associationwith Parduscoswere ConirostrumJerrugineiventre, C. sitticolor,Diglossa la/resnayii, Anisognathus igniventris, Dubusia cas- taneoventris,Hemispingus xanthophthalmus, H. ru/osuperciliaris,H. tri]asciatus,and Buthraupisaureodorsalis. Also observedin closeprox- imity to Parduscoswere Iridosornisjelskii, Ochthoeca]umicolor, Am- pelion [Doliornis]sclateri, Synallaxis gularis, and Schizoeaca]uliginosa. The only possibleavian predator recorded in the same woodlandswas an occasional Falco femoralis.

TAXONOMIC AFFINITIES

Although these behavioral notes are of considerableinterest because they concerna previouslyunknown species and one that still only three ornithologistshave seen alive, they provide no substantialclues to the bird's taxonomic affinities. Unfortunately, few of the conventional anatomical features employed in characterizing oscine families and subfamiliesare truly diagnosticfor all membersof a given group. For example, in the assemblageo.f genera presently constituting the Thraupinae, the presenceor absenceof rictal and facial bristles varies to almost every degree. In some genera (e.g. Habia) rictal bristles are numerous and pronounced,in other genera (e.g. Tangara) they are sparseand weakly developed,and in still other genera (e.g. Thraupis) they are obsoleteor absent. In the Emberizinae rictal bristles are sometimespresent and well developed (e.g. Pipilo) although usually barely noticeableo.r lacking (e.g. Sicalis). The same difficulty is evident with respect to other anatomical features such as the presenceor absenceof a nasal operculum. In the Emberizinae,for example, this structureis lackingin mostgenera but is presentin some(e.g. Aimophila). In the case of foraging habits, a set method is seldom diagnosticfor all membersof one subfamily. In most generaof the subfamily Car- duelinae,as now constituted,feeding is primarily arboreal, but in some July 1976] The Pardusco 427 generait is terrestrial. In short, in any one family or subfamily, not only do individual morphologicalcharacters run a gamut of variation but so do certain behavioral patterns. The allocation of Nephelornis is especially difficult becausethe bird is by no means readily assignableto one subfamily or another. None of the taxa to which it might conceivablybelong is characterizedby one or more clearly diagnosticanatomical features. The slightly frilled tongue with its shallow groove has remotely similar counterpartsin certain honeycreepers,but unfortunately the genera involved therein are themselvesof uncertain systematic affinities. Nephelornis oneilli and Xenodacnisparina seem to fill similar ecologicalniches and the two speciesare behaviorally somewhat alike. But Nephelornis oneilli feedson nectaronly secondarily,if at all, and in colorationit doesnot resembleXeno.dacnis parina, much less any speciesof Diglossa. Conse- quently,it is probablynot closelyrelated to either of thesegenera. ConceivablyNephelornis might be relegatedto the Thraupinae,for that assemblageas presentlyconstituted certainly containsmany diverse types, but its placementthere would merely add another anomaly to what is already an extremelyheterogeneous group that is possiblyeven now polyphyletic. Becausea study of the external morphologyleads to no conclusive decisionwith regard to the proper placementof Nephelornis,we must fall back on its unique tongue musclatureas perhaps the most trust- worthy index to its correct taxonomicallocation. As previously noted, it possessesone muscle that is lost in other nine-primaried oscines, and it has another musclewhose origin is different from that of the same musclein other nine-primariedoscines. The two musclessuggest that Nephelornisis a primitive offshootin the early radiation of the nine-primariedassemblage. Consequently, until new considerationsare brought to bear on the problem, we recommendthat Nephelornisbe listed next to Conirostrum among the genera incertae sedis that imme- diately follow the Parulinae. This position appears to be as near the base of the nine-primaried oscinesas our present classificationpermits Nephelornisto be placed (Lowery and Monroe 1968; Paynter 1970; Storer 1970, 1971, and in litt.). New avenuesof inquiry that involve biochemicalanalyses show promiseof eventually revealingnot only the true relationshipsof Nephelornisbut of other questionabletaxa as well.

ACKNOWLEDGMENTS

We continue to be grateful for the generousand unceasingsupport that John S. McIlhenny of Baton Rouge has given to the field explorations in Peru by the LSUMZ for more than a decade. He follows every development with the most heartwarming and enthusiastic interest. Likewise of great help was the munificent 428 LOWERYAND TALLMAI• [Auk, Vol. 93 contributionprovided by anotherpatron of the museum,John F. Maher, of Houston, Texas. To both of these gentlemenwe extend sincereappreciation, not only in our own behalf but also in behalf of Louisiana State University and its Museum of Zoology. We thank our associates,including Robert S. Kennedy, Gary Lester, Ronald J. Louque, Carol S. O'Neill, John P. O'Neill, Theodore Parker, III, Reyes Rivera A., Erika J. Tallman, and Manuel Villar, for their untiring efforts in making the field work of 1973-1975 successfulin ways that exceededall expectations.We also expressour appreciationto Marc Dourojeanni R., Carlos Ponce P., and Antonio Brack E. of the Direcci6n General Forestal y de Fauna, Ministerio de Agricultura, Lima, Peril, for their interest in our work and for the issuanceof scientific col- lecting permits. Dr. Hernando de Macedo R. of the Museo de Hlstoria Natural "Javier Prado," of Lima also rendered assistanceto us in many ways. Robert J. Newman offered valuable suggestionsand Walter J. Bock furnished indispensable help with anatomical matters and provided us with drawings made under his direc- tion by Dorothea Goldys.

LITERATURE CITED LOWERY,G. H., JR., ANDBURT L. MONROE,JR. 1968. Family Parulidae. Pp. 3-93 in Check-listof birds of the world, vol. 14 (R. A. Paynter, Jr., Ed.). Cambridge, Mass., Museum of Comparative Zoology. PA¾>•TEa,R. A., JR. 1970. Subfamily Emberizinae. Pp. 3-214 in Check-listof birds of the world, •ol. 13 (R. A. Paynter, Jr., Ed.). Cambridge,Mass., Museum of Comparative Zoology. RInGWAY,R. 1912. Color standards and color nomenclature. Washington, D.C., publishedby the author. STORER,R.W. 1970. Subfamily Thraupinae. Pp. 246-408 in Check-list of birds of the world, vol. 13 (R. A. Paynter, Jr., Ed.). Cambridge, Mass., Museum of Comparative Zoology. STORER,R. W. 1971. Classification of birds. Pp. 1-18 in Arian biology, vol. 1 (D. S. Farner and J. R. King, Eds.). New York• Academic Press. Z•MMER, J. T. 1930. Birds of the Marshall Field Peruvian expedition, 1922-1923. Field Mus. Nat. Hist. Zool. Ser. 17: 231-480.

Museumof Zoology,Louisiana State University,Baton Rouge,Louisi- ana 70893. Accepted12 February 1976. This paper was subsidizedby a friend of the L.S.U.M.Z.